Seabirds reveal mercury distribution across the North Atlantic.

Mercury (Hg) is a heterogeneously distributed toxicant affecting wildlife and human health. Yet, the spatial distribution of Hg remains poorly documented, especially in food webs, even though this knowledge is essential to assess large-scale risk of toxicity for the biota and human populations. Here, we used seabirds to assess, at an unprecedented population and geographic magnitude and high resolution, the spatial distribution of Hg in North Atlantic marine food webs. To this end, we combined tracking data of 837 seabirds from seven different species and 27 breeding colonies located across the North Atlantic and Atlantic Arctic together with Hg analyses in feathers representing individual seabird contamination based on their winter distribution. Our results highlight an east-west gradient in Hg concentrations with hot spots around southern Greenland and the east coast of Canada and a cold spot in the Barents and Kara Seas. We hypothesize that those gradients are influenced by eastern (Norwegian Atlantic Current and West Spitsbergen Current) and western (East Greenland Current) oceanic currents and melting of the Greenland Ice Sheet. By tracking spatial Hg contamination in marine ecosystems and through the identification of areas at risk of Hg toxicity, this study provides essential knowledge for international decisions about where the regulation of pollutants should be prioritized.

Spatial distribution of selenium-mercury in Arctic seabirds.

Mercury (Hg) is a metallic trace element toxic for humans and wildlife that can originate from natural and anthropic sources. Hg spatial gradients have been found in seabirds from the Arctic and other oceans, suggesting contrasting toxicity risks across regions. Selenium (Se) plays a protective role against Hg toxicity, but its spatial distribution has been much less investigated than that of Hg. From 2015 to 2017, we measured spatial co-exposure of Hg and Se in blood samples of two seabird species, the Brünnich's guillemot (Uria lomvia) and the black-legged kittiwake (Rissa tridactyla) from 17 colonies in the Arctic and subarctic regions, and we calculated their molar ratios (Se:Hg), as a measure of Hg sequestration by Se and, therefore, of Hg exposure risk. We also evaluated concentration differences between species and ocean basins (Pacific-Arctic and Atlantic-Arctic), and examined the influence of trophic ecology on Hg and Se concentrations using nitrogen and carbon stable isotopes. In the Atlantic-Arctic ocean, we found a negative west-to-east gradient of Hg and Se for guillemots, and a positive west-to-east gradient of Se for kittiwakes, suggesting that these species are better protected from Hg toxicity in the European Arctic. Differences in Se gradients between species suggest that they do not follow environmental Se spatial variations. This, together with the absence of a general pattern for isotopes influence on trace element concentrations, could be due to foraging ecology differences between species. In both oceans, the two species showed similar Hg concentrations, but guillemots showed lower Se concentrations and Se:Hg than kittiwakes, suggesting a higher Hg toxicity risk in guillemots. Within species, neither Hg, nor Se or Se:Hg differed between both oceans. Our study highlights the importance of considering Se together with Hg, along with different species and regions, when evaluating Hg toxic effects on marine predators in international monitoring programs.

Hybridization of Atlantic puffins in the Arctic coincides with 20th-century climate change.

The Arctic is experiencing the fastest rates of global warming, leading to shifts in the distribution of its biota and increasing the potential for hybridization. However, genomic evidence of recent hybridization events in the Arctic remains unexpectedly rare. Here, we use whole-genome sequencing of contemporary and 122-year-old historical specimens to investigate the origin of an Arctic hybrid population of Atlantic puffins (Fratercula arctica) on Bjørnøya, Norway. We show that the hybridization between the High Arctic, large-bodied subspecies F. a. naumanni and the temperate, smaller-sized subspecies F. a. arctica began as recently as six generations ago due to an unexpected southward range expansion of F. a. naumanni. Moreover, we find a significant temporal loss of genetic diversity across Arctic and temperate puffin populations. Our observations provide compelling genomic evidence of the impacts of recent distributional shifts and loss of diversity in Arctic communities during the 20th century.

Molecular tools prove little auks from Svalbard are extremely selective for Calanus glacialis even when exposed to Atlantification.

Two Calanus species, C. glacialis and C. finmarchicus, due to different life strategies and environmental preferences act as an ecological indicators of Arctic Atlantification. Their high lipid content makes them important food source for higher trophic levels of Arctic ecosystems including the most abundant Northern Hemisphere's seabird, the little auk (Alle alle). Recent studies indicate a critical need for the use of molecular methods to reliably identify these two sympatric Calanus species. We performed genetic and morphology-based identification of 2600 Calanus individuals collected in little auks foraging grounds and diet in summer seasons 2019-2021 in regions of Svalbard with varying levels of Atlantification. Genetic identification proved that 40% of Calanus individuals were wrongly classified as C. finmarchicus according to morphology-based identification in both types of samples. The diet of little auks consisted almost entirely of C. glacialis even in more Atlantified regions. Due to the substantial bias in morphology-based identification, we expect that the scale of the northern expansion of boreal C. finmarchicus may have been largely overestimated and that higher costs for birds exposed to Atlantification could be mostly driven by a decrease in the size of C. glacialis rather than by shift from C. glacialis to C. finmarchicus.

Ecological impacts of climate change on Arctic marine megafauna.

Global warming affects the Arctic more than any other region. Mass media constantly relay apocalyptic visions of climate change threatening Arctic wildlife, especially emblematic megafauna such as polar bears, whales, and seabirds. Yet, we are just beginning to understand such ecological impacts on marine megafauna at the scale of the Arctic. This knowledge is geographically and taxonomically biased, with striking deficiencies in the Russian Arctic and strong focus on exploited species such as cod. Beyond a synthesis of scientific advances in the past 5 years, we provide ten key questions to be addressed by future work and outline the requested methodology. This framework builds upon long-term Arctic monitoring inclusive of local communities whilst capitalising on high-tech and big data approaches.

Wind-driven upwelling of iron sustains dense blooms and food webs in the eastern Weddell Gyre.

The Southern Ocean is a major sink of anthropogenic CO2 and an important foraging area for top trophic level consumers. However, iron limitation sets an upper limit to primary productivity. Here we report on a considerably dense late summer phytoplankton bloom spanning 9000 km2 in the open ocean of the eastern Weddell Gyre. Over its 2.5 months duration, the bloom accumulated up to 20 g C m-2 of organic matter, which is unusually high for Southern Ocean open waters. We show that, over 1997-2019, this open ocean bloom was likely driven by anomalies in easterly winds that push sea ice southwards and favor the upwelling of Warm Deep Water enriched in hydrothermal iron and, possibly, other iron sources. This recurring open ocean bloom likely facilitates enhanced carbon export and sustains high standing stocks of Antarctic krill, supporting feeding hot spots for marine birds and baleen whales.

Extreme snowstorms lead to large-scale seabird breeding failures in Antarctica.

Climate change increases the frequency and intensity of extreme weather events that negatively impact wildlife, from individuals to whole ecosystems1. In polar environments, such events include heat waves2, anomalous sea ice concentrations3 and storms4. Polar seabirds are adapted to withstand harsh conditions, and although extreme weather events affect their breeding success and other demographic rates, they are thought to affect only a part of the population. Complete breeding failure of an entire population due to extreme environmental conditions is rarely observed5. Here we report how exceptional storm activity in Dronning Maud Land (DML), Antarctica, in the austral summer of 2021/2022 caused almost complete and large-scale breeding failures of the area's three most common seabird species - Antarctic petrel (Thalassoica antarctica), Snow petrel (Pagodroma nivea) and South polar skua (Stercorarius maccormicki).

Climate change in the Arctic: Testing the poleward expansion of ticks and tick-borne diseases.

Climate change is most strongly felt in the polar regions of the world, with significant impacts on the species that live there. The arrival of parasites and pathogens from more temperate areas may become a significant problem for these populations, but current observations of parasite presence often lack a historical reference of prior absence. Observations in the high Arctic of the seabird tick Ixodes uriae suggested that this species expanded poleward in the last two decades in relation to climate change. As this tick can have a direct impact on the breeding success of its seabird hosts and vectors several pathogens, including Lyme disease spirochaetes, understanding its invasion dynamics is essential for predicting its impact on polar seabird populations. Here, we use population genetic data and host serology to test the hypothesis that I. uriae recently expanded into Svalbard. Both black-legged kittiwakes (Rissa tridactyla) and thick-billed murres (Uria lomvia) were sampled for ticks and blood in Kongsfjorden, Spitsbergen. Ticks were genotyped using microsatellite markers and population genetic analyses were performed using data from 14 reference populations from across the tick's northern distribution. In contrast to predictions, the Spitsbergen population showed high genetic diversity and significant differentiation from reference populations, suggesting long-term isolation. Host serology also demonstrated a high exposure rate to Lyme disease spirochaetes (Bbsl). Targeted PCR and sequencing confirmed the presence of Borrelia garinii in a Spitsbergen tick, demonstrating the presence of Lyme disease bacteria in the high Arctic for the first time. Taken together, results contradict the notion that I. uriae has recently expanded into the high Arctic. Rather, this tick has likely been present for some time, maintaining relatively high population sizes and an endemic transmission cycle of Bbsl. Close future observations of population infestation/infection rates will now be necessary to relate epidemiological changes to ongoing climate modifications.

Gone with the wind - Wind speed affects prey accessibility for a High Arctic zooplanktivorous seabird, the little auk Alle alle.

Foraging ecology of chick rearing seabirds is affected mainly by the food availability on feeding grounds, but it can be also modulated by environmental conditions during the foraging trip, in that wind force. Considering predicted strengthening of surface winds over the Arctic Ocean, this factor may have a growing impact on the foraging performance of Arctic seabirds. Here, we studied how wind speed could affect prey accessibility for the High Arctic zooplanktivorous seabird, the little auk Alle alle breeding in Svalbard in 2015-2019. First, we estimated availability of its preferred prey, a cold water copepod Calanus glacialis, based on wider-scale mesozooplankton biomass model and environmental conditions. Then we estimated prey accessibility by including wind speed, the factor affecting the flapping flight performance of little auks commuting from/to the colony. Finally, we compared reproductive performance of the little auks (chick diet, growth rate and survival and duration of foraging flights of adults) between the studied years differing in wind and food availability conditions. We found that wind speed could affect significantly food accessibility for a zooplanktivorous seabird. Despite high spatial and temporal variability in prey availability and accessibility in shelf waters of SW Spitsbergen, interannual differences in duration of foraging flights and chick growth rate, little auks were able to sustain high breeding success confirming their capacity to buffer suboptimal foraging conditions. Our multidisciplinary work, combining multi-year remote sensing of oceanographic conditions, zooplankton availability and accessibility modelling, little auks diet composition and chick growth and survival emphasizes the importance of including wind conditions in the studies of foraging ecology of seabirds.

Foraging range scales with colony size in high-latitude seabirds.

Density-dependent prey depletion around breeding colonies has long been considered an important factor controlling the population dynamics of colonial animals.1-4 Ashmole proposed that as seabird colony size increases, intraspecific competition leads to declines in reproductive success, as breeding adults must spend more time and energy to find prey farther from the colony.1 Seabird colony size often varies over several orders of magnitude within the same species and can include millions of individuals per colony.5,6 As such, colony size likely plays an important role in determining the individual behavior of its members and how the colony interacts with the surrounding environment.6 Using tracking data from murres (Uria spp.), the world's most densely breeding seabirds, we show that the distribution of foraging-trip distances scales to colony size0.33 during the chick-rearing stage, consistent with Ashmole's halo theory.1,2 This pattern occurred across colonies varying in size over three orders of magnitude and distributed throughout the North Atlantic region. The strong relationship between colony size and foraging range means that the foraging areas of some colonial species can be estimated from colony sizes, which is more practical to measure over a large geographic scale. Two-thirds of the North Atlantic murre population breed at the 16 largest colonies; by extrapolating the predicted foraging ranges to sites without tracking data, we show that only two of these large colonies have significant coverage as marine protected areas. Our results are an important example of how theoretical models, in this case, Ashmole's version of central-place-foraging theory, can be applied to inform conservation and management in colonial breeding species.

Variation and correlation in the timing of breeding of North Atlantic seabirds across multiple scales.

Timing of breeding, an important driver of fitness in many populations, is widely studied in the context of global change, yet despite considerable efforts to identify environmental drivers of seabird nesting phenology, for most populations we lack evidence of strong drivers. Here we adopt an alternative approach, examining the degree to which different populations positively covary in their annual phenology to infer whether phenological responses to environmental drivers are likely to be (a) shared across species at a range of spatial scales, (b) shared across populations of a species or (c) idiosyncratic to populations. We combined 51 long-term datasets on breeding phenology spanning 50 years from nine seabird species across 29 North Atlantic sites and examined the extent to which different populations share early versus late breeding seasons depending on a hierarchy of spatial scales comprising breeding site, small-scale region, large-scale region and the whole North Atlantic. In about a third of cases, we found laying dates of populations of different species sharing the same breeding site or small-scale breeding region were positively correlated, which is consistent with the hypothesis that they share phenological responses to the same environmental conditions. In comparison, we found no evidence for positive phenological covariation among populations across species aggregated at larger spatial scales. In general, we found little evidence for positive phenological covariation between populations of a single species, and in many instances the inter-year variation specific to a population was substantial, consistent with each population responding idiosyncratically to local environmental conditions. Black-legged kittiwake Rissa tridactyla was the exception, with populations exhibiting positive covariation in laying dates that decayed with the distance between breeding sites, suggesting that populations may be responding to a similar driver. Our approach sheds light on the potential factors that may drive phenology in our study species, thus furthering our understanding of the scales at which different seabirds interact with interannual variation in their environment. We also identify additional systems and phenological questions to which our inferential approach could be applied.

Mercury contamination and potential health risks to Arctic seabirds and shorebirds.

Since the last Arctic Monitoring and Assessment Programme (AMAP) effort to review biological effects of mercury (Hg) on Arctic biota in 2011 and 2018, there has been a considerable number of new Arctic bird studies. This review article provides contemporary Hg exposure and potential health risk for 36 Arctic seabird and shorebird species, representing a larger portion of the Arctic than during previous AMAP assessments now also including parts of the Russian Arctic. To assess risk to birds, we used Hg toxicity benchmarks established for blood and converted to egg, liver, and feather tissues. Several Arctic seabird populations showed Hg concentrations that exceeded toxicity benchmarks, with 50 % of individual birds exceeding the "no adverse health effect" level. In particular, 5 % of all studied birds were considered to be at moderate or higher risk to Hg toxicity. However, most seabirds (95 %) were generally at lower risk to Hg toxicity. The highest Hg contamination was observed in seabirds breeding in the western Atlantic and Pacific Oceans. Most Arctic shorebirds exhibited low Hg concentrations, with approximately 45 % of individuals categorized at no risk, 2.5 % at high risk category, and no individual at severe risk. Although the majority Arctic-breeding seabirds and shorebirds appeared at lower risk to Hg toxicity, recent studies have reported deleterious effects of Hg on some pituitary hormones, genotoxicity, and reproductive performance. Adult survival appeared unaffected by Hg exposure, although long-term banding studies incorporating Hg are still limited. Although Hg contamination across the Arctic is considered low for most bird species, Hg in combination with other stressors, including other contaminants, diseases, parasites, and climate change, may still cause adverse effects. Future investigations on the global impact of Hg on Arctic birds should be conducted within a multi-stressor framework. This information helps to address Article 22 (Effectiveness Evaluation) of the Minamata Convention on Mercury as a global pollutant.

Fine-scale spatial segregation in a pelagic seabird driven by differential use of tidewater glacier fronts.

In colonially breeding marine predators, individual movements and colonial segregation are influenced by seascape characteristics. Tidewater glacier fronts are important features of the Arctic seascape and are often described as foraging hotspots. Albeit their documented importance for wildlife, little is known about their structuring effect on Arctic predator movements and space use. In this study, we tested the hypothesis that tidewater glacier fronts can influence marine bird foraging patterns and drive spatial segregation among adjacent colonies. We analysed movements of black-legged kittiwakes (Rissa tridactyla) in a glacial fjord by tracking breeding individuals from five colonies. Although breeding kittiwakes were observed to travel up to ca. 280 km from the colony, individuals were more likely to use glacier fronts located closer to their colony and rarely used glacier fronts located farther away than 18 km. Such variation in the use of glacier fronts created fine-scale spatial segregation among the four closest (ca. 7 km distance on average) kittiwake colonies. Overall, our results support the hypothesis that spatially predictable foraging patches like glacier fronts can have strong structuring effects on predator movements and can modulate the magnitude of intercolonial spatial segregation in central-place foragers.

Influence of reproductive output on divorce rates in polar seabirds.

The high occurrence of social monogamy in birds has led to questions about partner fidelity, or the perennial nature of monogamy from one breeding season to another. Despite the evolutionary advantages of partner fidelity, divorce occurs among 95% of bird species. We aimed to describe patterns of divorce and partner fidelity in five seabird species breeding in Arctic and Antarctic regions and investigated the influence of breeding status on pair bond maintenance. For four out of the five species considered, we observed low divorce rates (respectively 1.9%, 3.3%, 2.5%, and 0.0% for Brünnich's guillemot, glaucous gull, Antarctic petrel, and south polar skua), while the divorce rate was much higher (19.1%) for the black-legged kittiwake. For kittiwakes, the divorce rate was lower for pairs that managed to raise their chick to 15 days of age, while the effect of breeding success on divorce in the four other species could not be tested due to the rareness of divorce events. Our results emphasize the potentially large temporal (interannual) variations that should be taken into account in understanding divorce and partner fidelity in seabirds.

North Atlantic winter cyclones starve seabirds.

Each winter, the North Atlantic Ocean is the stage for numerous cyclones, the most severe ones leading to seabird mass-mortality events called "winter wrecks."1-3 During these, thousands of emaciated seabird carcasses are washed ashore along European and North American coasts. Winter cyclones can therefore shape seabird population dynamics4,5 by affecting survival rates as well as the body condition of surviving individuals and thus their future reproduction. However, most often the geographic origins of impacted seabirds and the causes of their deaths remain unclear.6 We performed the first ocean-basin scale assessment of cyclone exposure in a seabird community by coupling winter tracking data for ∼1,500 individuals of five key North Atlantic seabird species (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia, and Rissa tridactyla) and cyclone locations. We then explored the energetic consequences of different cyclonic conditions using a mechanistic bioenergetics model7 and tested the hypothesis that cyclones dramatically increase seabird energy requirements. We demonstrated that cyclones of high intensity impacted birds from all studied species and breeding colonies during winter but especially those aggregating in the Labrador Sea, the Davis Strait, the surroundings of Iceland, and the Barents Sea. Our broad-scale analyses suggested that cyclonic conditions do not increase seabird energy requirements, implying that they die because of the unavailability of their prey and/or their inability to feed during cyclones. Our study provides essential information on seabird cyclone exposure in a context of marked cyclone regime changes due to global warming.8.

Using near-infrared reflectance spectroscopy (NIRS) to estimate carbon and nitrogen stable isotope composition in animal tissues.

Stable isotopes analysis (SIA) of carbon and nitrogen provides valuable information about trophic interactions and animal feeding habits.We used near-infrared reflectance spectroscopy (NIRS) and support vector machines (SVM) to develop a model for screening isotopic ratios of carbon and nitrogen (δ 13C and δ 15N) in samples from living animals. We applied this method on dried blood samples from birds previously analyzed for δ 13C and δ 15N to test whether NIRS can be applied to accurately estimate isotopic ratios.Our results show a prediction accuracy of NIRS (R 2 > 0.65, RMSEP < 0.28) for both δ 13C and δ 15N, representing a 12% of the measurement range in this study.Our study suggests that NIRS can provide a time- and cost-efficient method to evaluate stable isotope ratios of carbon and nitrogen when substantial differences in δ 13C or δ 15N are expected, such as when discriminating among different trophic levels in diet.

Complex population structure of the Atlantic puffin revealed by whole genome analyses.

The factors underlying gene flow and genomic population structure in vagile seabirds are notoriously difficult to understand due to their complex ecology with diverse dispersal barriers and extensive periods at sea. Yet, such understanding is vital for conservation management of seabirds that are globally declining at alarming rates. Here, we elucidate the population structure of the Atlantic puffin (Fratercula arctica) by assembling its reference genome and analyzing genome-wide resequencing data of 72 individuals from 12 colonies. We identify four large, genetically distinct clusters, observe isolation-by-distance between colonies within these clusters, and obtain evidence for a secondary contact zone. These observations disagree with the current taxonomy, and show that a complex set of contemporary biotic factors impede gene flow over different spatial scales. Our results highlight the power of whole genome data to reveal unexpected population structure in vagile marine seabirds and its value for seabird taxonomy, evolution and conservation.

As the Arctic becomes boreal: ongoing shifts in a high-Arctic seabird community.

The Arctic is currently experiencing the most rapid warming on Earth. Arctic species communities are expected to be restructured with species adapted to warmer conditions spreading poleward and, if already present, becoming more abundant. We tested this prediction using long-term monitoring data (2009-2018) from nine of the most common seabird species breeding in the High Arctic Svalbard archipelago. This region is characterized by rapidly warming ocean temperatures, declining sea-ice concentrations and an increasing influence of Atlantic waters. Concurrent with these environmental changes, we found a shift in the Svalbard seabird community, with an increase in abundance of boreal species (defined here as species breeding commonly in temperate environments) and a decline in Arctic species (species breeding predominantly in the Arctic). Combined with previous observations from lower trophic levels, our results confirmed that part of the Arctic fauna is moving from an arctic to a boreal (or north temperate) state, a process referred to as a "borealization." Spatial variations exist among colonies for some species, indicating that local conditions may affect the trajectories of specific populations and potentially counterbalance the consequences of large-scale climate warming.

Within and between breeding-season changes in contaminant occurrence and body condition in the Antarctic breeding south polar skua.

The Antarctic ecosystem represents a remote region far from point sources of pollution. Still, Antarctic marine predators, such as seabirds, are exposed to organohalogen contaminants (OHCs) which may induce adverse health effects. With increasing restrictions and regulations on OHCs, the levels and exposure are expected to decrease over time. We studied south polar skua (Catharacta maccormiciki), a top predator seabird, to compare OHC concentrations measured in whole blood from 2001/2002 and 2013/2014 in Dronning Maud Land. As a previous study found increasing organochlorine concentrations with sampling day during the 2001/2002 breeding season, suggesting dietary changes, we investigated if this increase was repeated in the 2013/2014 breeding season. In addition to organochlorines, we analyzed hydroxy-metabolites, brominated contaminants and per- and polyfluoroalkyl substances (PFAS) in 2013/2014, as well as dietary descriptors of stable isotopes of carbon and nitrogen, to assess potential changes in diet during breeding. Lipid normalized concentrations of individual OHCs were 63%, 87% and 105% higher for hexachlorobenzene (HCB), 1,1-dichloro-2,2-bis (p-chlorophenyl)ethylene (p,p'-DDE), and ∑Polychlorinated biphenyls (PCBs), respectively, in 2013/2014 compared to 2001/2002. South polar skuas males in 2013/2014 were in poorer body condition than in 2001/2002, and with higher pollutant levels. Poorer body condition may cause the remobilization of contaminants from stored body reserves, and continued exposure to legacy contaminants at overwintering areas may explain the unexpected higher OHC concentrations in 2013/2014 than 2001/2002. Concentrations of protein-associated PFAS increased with sampling day during the 2013/2014 breeding season, whereas the lipid-soluble chlorinated pesticides, PCBs and polybrominated diphenyl ether (PBDEs) showed no change. OHC occurrence was not correlated with stable isotopes. The PFAS biomagnification through the local food web at the colony should be investigated further.

Centennial relationships between ocean temperature and Atlantic puffin production reveal shifting decennial trends.

The current warming of the oceans has been shown to have detrimental effects for a number of species. An understanding of the underlying mechanisms may be hampered by the non-linearity and non-stationarity of the relationships between temperature and demography, and by the insufficient length of available time series. Most demographic time series are too short to study the effects of climate on wildlife in the classical sense of meteorological patterns over at least 30 years. Here we present a harvest time series of Atlantic puffins (Fratercula arctica) that goes back as far as 1880. It originates in the world's largest puffin colony, in southwest Iceland, which has recently experienced a strong decline. By estimating an annual chick production index for 128 years, we found prolonged periods of strong correlations between local sea surface temperature (SST) and chick production. The sign of decennial correlations switches three times during this period, where the phases of strong negative correlations between puffin productivity and SST correspond to the early 20th century Arctic warming period and to the most recent decades. Most of the variation (72%) in chick production is explained by a model in which productivity peaks at an SST of 7.1°C, clearly rejecting the assumption of a linear relationship. There is also evidence supporting non-stationarity: The SST at which puffins production peaked has increased by 0.24°C during the 20th century, although the increase in average SST during the same period has been more than three times faster. The best supported models indicate that the population's decline is at least partially caused by the increasing SST around Iceland.